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Fireflies in the Snow: Observations on two early-season arboreal fireflies Ellychnia corrusca and Pyractomena borealis

  • Emory River Land Company and Great Smoky Mountains Conservation Association

Abstract and Figures

Field studies on two cold-hardy sympatric firefly species Ellychnia corrusca complex (Linnaeus) and Pyractomena borealis (Randall) were conducted from 2008 to 2011, in East Tennessee in the southeastern United States. Their behaviours , predation events, escape strategies, calendar and degree-day parameters for life events are presented and regional differences discussed. Adults of E. corrusca and last instar larvae of P. borealis become active in February when snow and below-freezing temperatures are common. They gather in the furrows on the warmer, sunlit south sides of still-leafless preferred trees, the majority of which are large diameter oaks (Quercus sp.) for E. corrusca, hickories (Carya sp.) for P. borealis and tulip poplars (Liriodendron tulipifera) for both firefly species. Ellychnia corrusca, lanternless and diurnal, abruptly appear low on colony trees in late winter (Jan-Feb), orienting to the sun and vertically to one another upon initial emergence. A 4-to 6-week quies-cent period follows this emergence during which time they remain on their colony tree slowly patrolling or resting head down in the bark furrows. Flight, dispersal and mating occur in mid-March to April. Diurnal P. borealis larvae climb from cold winter ground up preferred trees seeking sunny, protected pupation sites. Frequent moving is common in Jan-Feb as larvae select ideal arboreal sites. Pupation takes place in March with protandrous males attaching, pupating and eclosing before most females. Adult males use two mating strategies. Initially day active, they seek out and tend both larval and pupal females for up to two weeks and attempt copulation as soon as the guarded virgin female ecloses, bypassing courtship and flashing. In the nocturnal courtship display, P. borealis males flash in flight every ca. 2 s in April on fair-weather nights above 10-12°C, high in the forest canopy. Intense male competition , repeated mating and prolonged (1-to 8-day) copulation is common.
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Fireies in the snow: Observations on two
early-season arboreal reies Ellychnia
corrusca and Pyractomena borealis
L F
Emory River Land Company, 11828 Couch Mill Road, Knoxville, TN USA 37932.
ABSTRACT Field studies on two cold-hardy sympatric rey species Ellychnia
corrusca complex (Linnaeus) and Pyractomena borealis (Randall) were conducted
from 2008 to 2011, in East Tennessee in the southeastern United States. eir behav-
iours, predation events, escape strategies, calendar and degree-day parameters for life
events are presented and regional dierences discussed. Adults of E. corrusca and last
instar larvae of P. borealis become active in February when snow and below-freezing
temperatures are common. ey gather in the furrows on the warmer, sunlit south
sides of still-leaess preferred trees, the majority of which are large diameter oaks
(Quercus sp.) for E. corrusca, hickories (Carya sp.) for P. borealis and tulip poplars
(Liriodendron tulipifera) for both rey species. Ellychnia corrusca, lanternless and
diurnal, abruptly appear low on colony trees in late winter (Jan-Feb), orienting to
the sun and vertically to one another upon initial emergence. A 4- to 6-week quies-
cent period follows this emergence during which time they remain on their colony
tree slowly patrolling or resting head down in the bark furrows. Flight, dispersal
and mating occur in mid-March to April. Diurnal P. borealis larvae climb from cold
winter ground up preferred trees seeking sunny, protected pupation sites. Frequent
moving is common in Jan-Feb as larvae select ideal arboreal sites. Pupation takes
place in March with protandrous males attaching, pupating and eclosing before most
females. Adult males use two mating strategies. Initially day active, they seek out and
tend both larval and pupal females for up to two weeks and attempt copulation as
soon as the guarded virgin female ecloses, bypassing courtship and ashing. In the
nocturnal courtship display, P. borealis males ash in ight every ca. 2 s in April on
fair-weather nights above 10-12°C, high in the forest canopy. Intense male competi-
tion, repeated mating and prolonged (1- to 8-day) copulation is common.
Keywords: Pyractomena borealis, Ellychnia corrusca, daytime lightning bug, mate
guarding, pupal tending, rey, preferred trees, arboreal, aerial pupation
Lampyrid 2012. 2:4 8-71
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Lampyrid Volume 2 (2012)
In early to mid-February (late winter)
with few insects evident, two sympatric
rey species become active in the bare,
cold, oen snow-covered forests of the
Great Tennessee (TN) Valley, the Great
Smoky Mountains and the Cumberland
Mountains of East TN in the south-
eastern United States (Fig. 1). Adults
of the Ellychnia corrusca L. complex
(Fender 1970), diurnal and lanternless,
emerge and share many of the same trees
as the last instar larvae of Pyractomena
borealis Randall that climb from the
forest oor to seek arboreal pupation sites on the southern aspect of the trunks of
those trees (Fig. 2).
Both species are widespread in the eastern half of the US with the latitudinal
range extending south from southeastern Canada to north central Florida (FL), a
distance of more than 2200 km (Green 1957; Fender 1970; Bousquet 1991; Lloyd
1997; Rooney & Lewis 2000) (Fig. 1). Initial natural history observations were
recorded for both species almost a century ago by Williams (1917). Pyractomena
borealis preimaginal larval descriptions are reported by Archangelsky and Branham
Figure 1 East TN sites (white circle: TN).
North-south range (red arrow) of both
Pyractomena borealis & Ellychnia corrusca
Figure 2 Ellychnia corrusca (A) and Pyractomena borealis (B). Note the pale elytral margin of
the latter.
Lampyrid Volume 2 (2012)
in 1998. Buschman (1984) describes the transformation of the prepupal pygypod
(moveable tail organ) to a permanent anchor-like organ in the closely related P.
lucifera. Aerial, arboreal pupation behaviours, such as tree positioning to inu-
ence microhabitats (Lloyd 1997), pupal development rates and protandry (Gentry
2003) are provided for the semi-tropical FL P. borealis (877 km south of TN). Cooler
Massachusetts climate (1324 kilometers north of TN) E. corrusca adult wintering
and mating behaviours and survival rates are described by Rooney and Lewis
(2000). Like many temperate region reies, E. corrusca and P. borealis have well-
dened seasons of appearance and peak activity (Faust 2004, Faust & Weston 2009).
In East TN, adults of E. corrusca appear in Jan-Feb and are rarely seen aer May;
the last instar larvae of P. borealis also appear in February with few adults seen aer
April. Local species representatives can be found in the Great Smoky Mountains
National Park collection (GSMNP), Gatlinburg, TN (Mayor 2006).
Active in the daylight and oen found below 3 m on tree trunks, the adults of E.
corrusca and larvae of P. borealis are easily accessible for live studies in their natural
habitat before the other common fourteen-plus East TN species (Faust & Weston
2009) emerge. Both rey species appear to return to the same or nearby trees
each season. ese reies, widespread but less abundant than other local rey
species, are found in loose colonies on the margins or within the secondary, mixed
hardwood forests of East TN, which are predominantly Appalachian oak-hickory
type (Fig. 3).
is longitudinal study, spanning four years of focused studies and 19 years of
eld observations, documents the continuity and stability of these populations and
presents emerging threats, particularly to the Knox County population which was
the primary study site.
e study site habitats shared by both species are described together. e species
details are dealt with separately. Regional dierences and similarities are discussed
and new information presented on selection behaviour (of host trees and direc-
tional orientation), aspects of development (life event dates, mortality factors and
survival rates) and reproduction (sex ratio, larval/pupal seeking and tending, mate
guarding, ashing and mating). Average calendar dates and modied growing
degree day values (mGDDC) for life stage landmarks (Faust & Weston 2009) are
oered when signicant as a reference point for future comparisons to far distant
populations. is paper was presented at the August 2010 Malaysian International
Firey Symposium. New data from the 20112012 season has been added to augment
the results reported at that meeting. It is hoped this information will further the
understanding of these adaptable early season species.
Study sites
ere were ve study sites in East TN, USA, i.e. Knox (35°56’38.13”N 84°12’15.97”W,
Lampyrid Volume 2 (2012)
274-305 m a.s.l.), Blount (35°36’55.05”N 83°40’07.32”W, 640 m, GSMNP), Sevier
(35°39’02.79”N 83°34’32.33”W, 731 m, GSMNP), Jeerson-Grainger (36°06’23.64”N
83°37’58.11”W, 305 m) and Morgan (36°12’11.52”N 84°35’17.14”W, 407 m). Fireies
of both species have been found at these sites every year since I started observing
them in 1992. Knox County, the primary site, was 46–66 km from the other four
sites. e Knox site consisted of a 4.3 km circular loop covering 15 ha of maturing
species-diverse (i.e., more than 111 species) Appalachian oak-hickory type secondary
forest (Oswalt 2007) that borders privately and publicly owned lands (Fig. 3). e
four secondary sites are generally similar in habitat and forest composition, but the
two GSMNP sites are higher in elevation (Faust 2010) and the Jeerson-Grainger
site includes an alluvial oodplain forest. Using ve sites enabled comparing the
reies’ tree usage and emergence patterns in slightly varying habitats within the
same region. Within the sites, Ellychnia corrusca colonies have been found at the
same locations every year and while there may be slight shis in trees used, the
number of trees occupied is fairly consistent. At Knox, the main colony occupies 29
trees located on a 0.5 ha karst hilltop rim while a smaller colony is found on 2 trees
0.5 km away. e circular 0.37 ha Blount site has a large colony occupying 20 trees
and a smaller colony on 6 trees 1 km away.
Field sampling
e studies for the two rey species took place over four seasons (2008–11). Outside
the rey season from May–January, monthly site visits recorded sightings of adults
or larvae on trees or the ground. e sites were visited weekly from the rst appear-
Figure 3 Preferred trees (9 species) utilised by Ellychnia corrusca and P yractomena b orealis in
relation to the composition of a typical Appalachian oak-hickory forest with ≥111 species.
Lampyrid Volume 2 (2012)
ance of E. corrusca, in Jan or Feb through early May (last sightings for P. borealis).
During the rst few surveys for each rey species, the season’s colony trees were
identied and marked. Colony trees of previous years were checked rst, followed
by neighbouring trees. In the rst year of this study, P. borealis exuviae persisting on
trees used in 2007 identied previous colony trees for that rey species. Searches
for additional colony trees within the sites were carried out each season. For all
colony trees, the following were recorded: species, diameter at breast height (dbh),
height, vertical position in canopy layer (canopy or understory), bark characteristics
(smooth, scaly, rough or furrowed; if furrowed, depth of furrow measured), and
location in relation to amount of solar radiation received (sunny or shaded).
Surveys at secondary sites were not continued if fewer than ve individuals were
found during the rst weeks of emergence for that particular species. In addition
to the weekly site visits, the primary site (Knox) was visited 1–3 times daily from
the week of rst appearance. For both rey species, site visits recorded the number
of individuals found on each of their respective colony trees. For each individual,
the following was also recorded: height from the ground; orientation in relation to:
1) aspect on tree (i.e., amount of solar radiation received, where the south-facing
side is warmest),
2) the vertical line of the tree trunk,
3) other individuals; activity level (slow or fast-moving); sex (of adults for E.
corrusca, when possible and at the pupal/adult stage for P. borealis). Aspects
of behaviour during primary life events were recorded (quiescence, escape
strategies, dispersal, mating and oviposition for E. corrusca; escape strate-
gies, pupation, pupal-guarding, eclosion, courtship and mating for P. borealis).
Mortality and likely causes were recorded, and also the presence of other insects
resembling these reies.
As P. borealis courtship displays commenced at Knox, nightly walks along the
loop trail were made to record the relative number of individuals in the treetops.
Individuals could be reliably counted by their ashes because they usually ashed
more than 5 m apart from their nearest closest neighbours. Towards the end of
the courtship period as the trees leafed out (thus obstructing canopy view from
beneath), ash counts were made from the creek bordering half the loop.
Additionally, in 2011 at Knox and Blount, all E. corrusca sighted were marked on
their elytrae with distinct individual patterns of colour/dots/dashes with non-toxic
ink (Uchida DecoColour™) identifying them by colony tree on the date they were
rst sighted. Daily (visual) recaptures were carried out to determine their movement
(if any) to other trees. Live elytral length was measured using a Mayo™ caliper and
ruler. Also in 2011 at Knox, about half of all P. borealis larvae sighted and later, all
eclosing adults, were likewise marked and recaptured (visually) daily. Over the 4
years, random live measurements of larvae, pupae and adults were made to deter-
mine general live size ranges.
Lampyrid Volume 2 (2012)
Accumulated modied growing degree-day (mGDD) values, also known as
30/10 C (or 86/50F corn growing degree days in US) and daily temperature data,
were obtained from National Weather Service (NWS) station #406742 for all study
sites except the higher elevation Sevier and Blount sites which used NWS #316341.
For more complete explanation of this heuristic tool as it applies to reies and
data tables see Faust and Weston (2009) and Fridley (2009), and the regional and
federal climate centre microclimates websites (NRCC 2008-2010; NOAA 2008–10).
Air temperatures were measured frequently and randomly comparing the microcli-
mates of the south- and north-facing sides of colony and preferred trees.
Laboratory and controlled eld studies
In 2010, E. corrusca adults (ve males and ve females) and a larva (a rare nd) were
observed in captivity for a week in April and September–October, respectively. e
adults were placed in a mesh-capped two-litre glass jar lined with native soil and a
few twigs positioned vertically. e larva was found glowing in the soil below leaf
litter by a gravel road at Sevier at 9 pm, 23rd September.
In 2008 and 2009, separate mesh enclosures were put around four P. borealis
female pupae where they were attached to their colony trees in order to determine
when they rst spontaneously ashed at males or articial ashes aer eclosion. Two
captive virgin females were observed in 2008 and 2009 for copulatory behaviour
and times. In 2011, 8 female and 5 male pupae were collected within several days
of predicted eclosion and placed in natural position, pygypodia taped to sticks, in
1 quart mesh covered containers in a covered outdoor setting, within their habitat.
Once eclosed, these 13 virgin adults were observed for 14 days for copulation events
and times and remating behaviours and times. Two additional wild caught males
were added to the remating studies the nal week. Apple slices were provided as
food. Aer 2-3 matings, females were returned to their pupal trees in order to
preserve the local population. Males were either released at night or kept until death
to determine typical male lifespan. Representative specimens of E. corrusca and P.
borealis from this study are deposited in the Florida State Collection of Arthro-
pods, Gainesville, FL. Photographs were taken with a Kodak Z740 Easyshare or an
Olympus Stylus 720 and 770SW.
Life events of Ellychnia corrusca
Like many diurnal reies with no lanterns, E. corrusca lacked paler elytral margins,
yet retain the bright red and pale yellow markings on their wide (3-7 mm) prono-
tums (Fig. 2). e ridged dorsal elytral surface was densely covered by yellowish,
olive green hairs. In the eld, both sexes looked similar. Size and pronotum colour
Lampyrid Volume 2 (2012)
variation was common. e body and elytral
length of specimens from Knox (16.2 mm ±
1.3 and 11.9 ± 1.1 mm, respectively; n = 19)
were greater than the higher elevation (731
m, 640 m) Sevier and Blount adults (12.9 ±
1.7 mm and 9.4 ± 1.4 mm, respectively; n =
44). Females from 9 - 17 mm in size were
seen coupled to males 9 mm in size (Fig. 4).
In 2010, three of 164 newly emergent adults
had bent or damaged elytra.
Robust, dark bodied Ellychnia corrusca
larvae, rarely seen, were covered in bristly
coarse setae. One glowing larva, found in
September, white and pink from recent
molting, had darkened by the next morning
(Fig. 5). It measured 15–17 mm long and
3.5 mm wide and readily ate small chopped
live worms 6 times in 21 days. It used its
pygypodia (caudal tail organ) for grooming
and to pry open a fresh water snail operculum. It did not attack a more fragile
Photinus sp. larva kept in the same enclosure. ey oen fed together, though the
Photinus larva deferred to it. Aer six weeks, it died from unknown causes.
Abrupt appearance and vertical emergence of adults
From late October to early February, few to no adults or larvae were evident either
on the ground or tree trunks. Adults were rst sighted at one or more of the sites
on 14, 10 and 18 February in
2008, 2009 and 2010, respec-
tively, and on 20 January in
2011. At Knox (29 trees) in
2010 (Fig. 6), no adults were
seen until 20 February when
44 were found. e next day’s
total was 104. Smaller groups
emerged over the nal week
in February, usually on clear
sunny days, with a late group of
seven appearing on 29 March
bringing to a total of 217 adults
for that site and 299 for all sites.
Figure 4 Ellychnia corrusca complex has
large size variation. Coupling occurs on
trees by day.
Figure 5 Ellychnia corrusca late instar larva feeding on a
fresh earthworm section.
Lampyrid Volume 2 (2012)
When emerging at temperatures below 5°C, oen with snow-covered ground, the
adults moved slowly. Mean height up the tree trunks increased daily from 66 cm (on
18 February) to 88 cm, 152 cm, and 170 cm on Feb 21 (range 15-152 cm, 22-243 cm,
96-307 cm, 0-320 cm respectively) at the Morgan and Knox sites with temperatures
at appearance ranging from 0-16°C.
ey were oriented facing a southerly direction (mean 182 ± 21°, n = 299) on
the trees, and on each tree were vertically oriented to one another (Fig. 7). In 2010,
at Knox, over 95% of 217 adults followed this
vertical orientation to one another. e rest
appeared scattered but still climbing the south-
erly aspect of the trees. On subsequent days, the
vertical arrangement disappeared as the majority
of adults scattered and climbed out of sight up
the trees. In 2011, at Knox and Blount, 90% of
the nal tally of adults (n = 123) emerged more
gradually from 29 January – 1 March with peak
numbers reached by mid-March. Orientation
was 189 ± 19°.
e 2012 season had an unusually mild winter
and early spring. First E. corrusca emergence of
30 climbing adults was observed on 29 December
2011 at the Blount site, with subsequent dates
of dispersal and coupling beginning 2-4 weeks
earlier than previously recorded (by 19 March).
Colony trees
From 2008-10, oaks (Quercus velutina, Q. alba
and Q. coccinea) and tulip poplars (Tulipifera
Figure 6 Visible presence of Ellychnia corrusca adults monthly for one year at Knox Co. colony,
in ight from July 2009 - June 2010 and on trees from July 2009 - April 2010.
Figure 7 Ellychnia corrusca, six
adults circled in black, abrupt
appearance in February with group
vertical tracking up the colony tree.
Lampyrid Volume 2 (2012)
liriodendron) were colonised by 87% (n = 241) of adults. ese tree species comprise
only 16% of the total forest composition of trees >2.5cm dbh (Oswalt 2007). At
Knox, in 2010 and 2011, 83% (n= 29) and 80% (n = 10) of the colony trees, respec-
tively, were oaks while at Blount, 84% (n = 26) of trees colonised were tulip poplars,
which are abundant there. Colony trees, 60 to 80 years old, were mature, living
and dominant canopy trees found in open deciduous forests. ey shared direct
exposure to winter sun for most of the day, had rough, furrowed or scaly bark
(furrow depth 12 ± 3.6 cm, n = 27), were canopy (25m+) trees and relatively large
dbh (57 ± 16 cm) compared to the many other tree species in the surrounding forest.
From 2008-2011, lightning, high winds and disease killed 4 of the colony trees.
e quiescent phase was from rst appearance to mid-March (Table 1). e adults
oen rested head-down in south facing bark furrows, with no apparent vertical
orientation to one another. ey were minimally active (crawling) on any sunny
above-freezing day. Marked adults (n = 123) in the recapture study frequently moved
up and down the trees, as much as 3 meters in 30 minutes especially on sunny days.
Up to 24 days aer the rst sighting, 96% of the 54 recaptured adults were on their
original tree. e remainder were on nearby trees (< 3 m away), having dropped to
the ground and disappeared when initially marked (which threatens them). In the
nal 2-3 weeks of quiescence before dispersal, no recaptures were recorded at Knox
and fewer than twelve were sighted at Blount. ey may have climbed up into the
colony tree canopies. Few were seen in late quiescence until ight, dispersal and
coupling began. In 2012, adults were visually recaptured as long as 66 days past
initial marking.
Dispersal, mating and ovipositing
Dispersal, recognised by the reappearance of reies in ight or active on both
colony and non-colony trees, occurred in mid-March at air temperatures above
15°C. First observed ight in 2010 and 2011 was on 8 March at 20°C and 17 March
at 23°C, respectively. On very sunny days where south-facing trunk temperatures
1 Early June for high elevation.
Table 1 Ellychnia corrusca general life cycle calendar, East Tennessee.
Larva e/
occass ional
Few or no
Emerge &
climb tr ee
Quiesce nt on
tree tru nk
Flight /
Couplin g /
Adult / deat h
May -
June -
February -
early March
February -
mid March
Mid - late
April Late April -
early May1
Lampyrid Volume 2 (2012)
exceeded 25°C, adults were oen seen on the north sides of the still-leaess trees
where temperatures were 5-10°C cooler. From 2008-11, coupling started in April
within 119-222 mGDDC (215-400 mGDDF), regardless of warm weather before
that time. At Knox and Sevier, the most active coupling period in 2010 and 2011 was
9-10 April (10 couples and 11 singles recorded) and 29 March – 9 April (11 couples
and 49 new unmarked singles recorded), respectively. In 2011, 23 and 19 colony and
non-colony trees, respectively, were used for courtship and coupling. Mating was
not observed on the ground. Flying individuals were most oen seen aer 12 noon
when it was warmest. By April’s end, few remained and no coupling was observed
at Knox while at Sevier, small numbers (<10) persisted through the rst week of June
in 2009, 2010, and 2011. From June-Dec, adults were rarely seen (Table 1 and Fig. 6).
In the laboratory, multiple matings between the 5 males and 5 females were
observed for both sexes, in Stage 1 and 2 positions (see Lewis & Wang 1991). Coupling
(9 observed in 7 days) took place vertically on twigs when available (otherwise on
the soil), commencing before sunrise and lasting 2 to more than 6 h. Females laid
o-white eggs measuring 0.5-0.9 mm (n= 27) singly in the soil on successive days.
Under magnication, the egg surface appeared slightly rough and sticky.
Escape strategies
When observed too closely or feeling threatened, their two primary escape strate-
gies were actively crawling up the trunk or dropping to the ground on their backs,
becoming well-camouaged in the leaf litter. Escape climbing took place even on
cold (<0°C) days, albeit slower than on warmer days (especially >20°C). When
marked, they exhibited distress by dropping to the ground, opening and closing
their elytra or attempting to rub their bodies on the bark. When handled or when
crawling, they oen deposited a clear droplet of liquid from their terminal segment.
ey exuded white beads of reexive bleeding from the elytral margins when
severely threatened. Flying, observed only aer dispersal, was seldom used as an
escape strategy.
Figure 8 Left to right: A white phorid (Apocephalus antennatus) larva emerges as its female
Ellychnia corrusca host dies; phorid pupae; adult phorid. Phorid ies are parasitoids of E.
corrusca and Pyractomena borealis.
Lampyrid Volume 2 (2012)
Predation and mimics
Predation was recorded early (28 February) in the season for 2010 and 2011 and
parasitism late (22 April) in the 2011 season. In 2010, two of four males taking
shelter in a hole at the base of a Quercus alba tree were preyed on and in 2011,
three unmarked elytra were found at the base of colony trees in thick leaf litter. Bite
marks and/or tears on elytral remnants were consistent with rodent or bird preda-
tion. In 2011, 32 phorid larvae (Brown 1994) that emerged from a female that died
shortly aer, were later identied (Brown, pers. comm.) as Apocephalus antennatus
Malloch (Diptera; Fig. 8). Beetles and moths from several families with mimic-like
colouration and patterns were oen found on or near the rey colony trees (Fig. 9).
Figure 9 Mimicry complex on/near trees colonised by Ellychnia corrusca and Pyractomena
borealis: 1-5. Cantharidae (8-12 mm); 6. Lycidae (9 mm); 7. Meloidae (6 mm); 8. Lampyridae (E.
corrusca on top, 15 mm; P. borealis beneath, 17 mm); 9. Carabidae (15 mm); 10. Pyrochoidae (12
mm); 11. Silphidae (20 mm); 12. Cerambycidae (9 mm); 13-14. Arctidae (20 mm); 15-17. Elateridae
(10 mm); 18. Pedilidae (7 mm).
Lampyrid Volume 2 (2012)
Life events of Pyractomena borealis
Fall larvae
Over the four years, a total of ve late instar larvae were observed in September and
October (outside the P. borealis season). ey were found glowing intermittently on
leaf litter and vegetation up to 20 cm from the ground.
Climbing-searching larvae
e rst sightings of last instar, searching-climbing larvae occurred during the day
on 1st March, 10th, 20th and 21st February in 2008-11, at temperatures of 12°C,
10°C, 16°C and 19°C, respectively. In the very mild winter of 2012, larvae climbed
over a month earlier (Jan 22), rst ashed and fully eclosed up to 3 weeks earlier
(19-25 Mar) than previous years. Male larvae climbed-searched 15 ± 8 d (n = 68,
2008-11) earlier than the females (n = 40). e pink-tinged larvae measured 16-26
mm (live length, n = 20). ey were narrow-headed, cryptically coloured, mottled
light yellow to dark brown. ey climbed up from the leaf litter and actively investi-
gated the south-facing furrows low on the trunks of their chosen trees. ey got no
higher than 1 m up the trees in the earliest searching phase in February and oen
disappeared early in the season. On very cold days (< -8°C), they remained motion-
less in the furrows. ey relocated every few days in search of more southerly or
protected sites.
By early March they were head down with dorsum facing outward, and had
begun to anchor their pygypodia within the furrows or under bark scales (Figs. 10
& 11). Four pupated ‘backwards’ with the dorsum facing the tree and/or (exposed)
outside the furrow. Anchoring was oen permanent within 3 days. Larvae disturbed
in the rst two days were found to abandon the site (12 of 21 larvae moved when
marked in 2011) instead of going on to pupate. Once attached, if disturbed, they
glowed and arched backwards and up.
e pupation sites were at a mean height of 147 ± 46 cm (n = 107) and oriented
at 170 ± 23°. In 4 years, the lowest pupation site was 28 cm and the highest observed
was 300 cm. Males pupated 12 ± 7 d (n = 68, 2008-11) earlier than females (n =
40) but their pupation heights appeared no dierent. Mean furrow depth for these
pupation sites was 10 ± 3 mm (n = 53).
Relative temperature measurements taken randomly at the actual pupation sites
throughout the rey season averaged 16°C (n = 320) compared to the opposite,
shadier side of the trees, which averaged 14°C. On sunny days before the trees leafed
out, the temperature dierence between the north- and south-facing pupation sites
oen exceeded 13°C.
During the larval climb and pupation period, temperatures ranged from -1°C to
31°C. e range for pronotal width was 4-7 mm and body length was 11-26 mm (n
= 45), with the largest larvae and pupae (22-26 mm, n = 21) eclosing into the largest
female adults (20-22 mm).
Lampyrid Volume 2 (2012)
By early April most of the larvae had pupated into the characteristic black pupae
with larval exuviae still attached to the pupal dorsum (Fig. 10). Two pupae at
pupation sites that reached extraordinarily high temperatures of 42°C (on 17 April
2010) failed to eclose. e two pupae that pupated backwards and outside a furrow
did not eclose but the two that pupated in exposed locations did eclose.
Males began to eclose late in March, 9 ± 7 days (n = 68, 2008-11) earlier than the
females (n = 40). e sexes could be dierentiated by their lanterns (Fig. 12). On cold
days with inclement weather (< 5°C), newly-eclosed adults stayed next to their pupal
exuviae for up to 24 h as they hardened and darkened while on hot days (> 25°C),
they quickly moved to a shadier location on the trunk.
Among the marked larvae in 2011, males and females began to eclose on 27
March and 3 April, respectively. By 8 April over 50% of the females (n = 15) had
eclosed, with the last male and female eclosing 9 April and 14 April respectively,
bringing to a total of 35 males and 15 females (Fig. 13). All were re-marked soon
aer eclosing. Eight males disappeared aer marking, but the other males remained
on their pupation trees. ere was no ashing from these rst-eclosing, day-active
searching males for over a week. ese males, diurnal at this stage, methodically
searched the trunks for larval or pupal females by day and rested motionless in
furrows at night (Figs. 10 & 11). Every pupa, furrow or scale 3 m up the trunk was
investigated. Upon locating a female, the male clasped her with his legs, pressed his
Larva Pu pa Adult eclos ing guarding pupa C oupling
Figure 10 Life stages of Pyractomena borealis. Protandrous males guard female larvae and
pupae and attempt coupling as female ecloses.
Lampyrid Volume 2 (2012)
pronotum to hers and became motionless. Pupae, when approached by a potential
guard male, arched back and forward rapidly (12 times in 2 min in one instance)
before being clasped and subdued by the male. A few male pupae (2 in 2011) were
guarded, but only briey (0.25-3.00 h). All pupae guarded for over a day eclosed
as females (2008-11). Competing males piled on top, grappled with, and oen
displaced the guard male, which would then be forced to search for another female
pupa. Males also used their pronotum to pry under or push away other males.
is competition intensied as females approached eclosion, with frequent power
reversals occurring among the increasing numbers of males. Female pupae 24 h
from eclosing had 3-7 males competing for them (n = 9).
From 2008-11, 79.1% (n = 43) of larval and pupal females were guarded by one or
more males from 1-14 days until eclosion (Figs. 10 & 11), which took about 30 min.
From 2008-11, 74.6% (n = 150) of eclosing adults were males and this male-biased
sex ratio was signicant (p<0.0001, exact binomial test). Preliminary molecular
studies on this local population found no evidence of Wolbachia bacteria (Werren
et al. 1995) as inuencing this ratio (F. Vencl, pers. comm. 2011) as has been found
to occur in some Korean rey species (Jeong et al. 2009).
Adult live mean body length, elytral length and pronotal width was 17 ± 2 mm,
11.5 ± 1.4 mm and 4.5 ± 1.2 mm, respectively (n = 58). Females, with distended
abdomens protruding 2-4 mm beyond the elytrae, oen measured longer than
males (20.1 ± 0.9 mm, n = 9 vs. 15.8 ± 1.7 mm, n = 35). Elytral length was more
similar: males 11 ± 1.2 mm; females 12 ± 1.6 mm (Fig. 12). In 2011, three of 88 newly
eclosed adults had deformed or missing elytrae and/or wings. is amount of defor-
mation was representative of previous years.
Figure 11 Pyractomena borealis males tend female larvae (A & B) and pupae up to two weeks
before eclosion. Disturbed males may abandon the female, but return when threat is gone.
Two and four males (C & D), some marked, grapple over a female pupa.
Lampyrid Volume 2 (2012)
Display and mating
Typically, a newly-eclosed (still
white and so) female (n = 18),
would be mobbed by her guard
and competing males. She would
rapidly leave her pupation site,
seeking cover under a bark scale
or crevice, with 1-3 males riding
her back. Copulation commenced
before her elytra had hardened and
darkened and no ash dialogue was
involved (Figs. 10 & 11).
From 2008-2011 rst ight and
ashing started on 10th, 12th, 11th
and 6th April respectively aer half the females had eclosed and climbed up the tree
disappearing in the canopy. In 2011, males then ceased patrolling the trunks and
even stopped guarding remaining pupae, decreasing from 46 guard males to 2 by 12
April with one female pupa le. Males displayed high in the bare or newly leafed-
out treetops, from 30 min aer sunset till sunrise with peak display 45-90 min aer
sunset. Temperatures were 14-22°C during display times. Peak nights recorded
50-120 observed individuals along the 4.3 km trail, showing that this species,
though common in dark forest habitats, is not abundant. ree ash patterns were
observed. e most common was the male’s single 0.3-0.5 s ash every 2-3 s. ey
ashed as they ew slowly around the canopy margins. Less common was the ash
pattern similar to the rst, but with the lantern dimly lit in between ashes. e
Figure 12 Lantern of male and female Pyractomena
borealis. Two lanterns on the male and four smaller
lanterns visible on the female.
Figure 13 Pyractomena borealis male and female eclosion, pupal guarding and display dates
for 2011. Note cool spell <10°C, with little activity, typical for this time of year, between 24
March and 2 April.
Lampyrid Volume 2 (2012)
rare female seen ashing (n=7, in the wild and in captivity) responded immediately
aer the male ash with a single or bimodal ash lasting ca. 0.70-1 s, with the initial
ash brighter.
In the controlled eld study, the 4 newly eclosed virgin females did not respond
to articial or male ashes until the third night aer eclosion. In the 2011 mating/
remating studies, the 8 virgin females were not initially receptive to the advances of
males introduced to the cage. Females ed, squeezed into crevices and tucked their
tails under their bodies when pursued. Despite this avoidance behaviour by the
females, males ultimately copulated successfully in 14 of the 16 observed matings
in captivity, but less oen in the wild where the female oen escaped the pursuing
male (unquantied pers. obs.) Copulations from 2008-2011, usually remaining in
Stage 1 position (Lewis & Wang 1991), lasted an average of 67 ± 64 h (n=17) (range
3-214 h), with mean virgin pairings lasting 66 ± 62 h (n=11) and mean re-matings
lasting 84 ± 74 h (n=5). Males stayed in copula even when probed or misted with
water. Once copulation began, competing males le. If a new male arrived, brief
skirmishes occurred as he tried to take the female, but did not succeed. Both males
and females re-mated readily (n= 8). Aer copulating, the males glowed continu-
ously. When released at the base of their colony trees, they continued glowing, but
aer 5 min switched to the characteristic ash pattern, all the while climbing up the
trees. In the controlled eld study, the rst male to eclose guarded 2 females for 9
days, coupled for 65 h in captivity with a third and died on the 24th day. One of the
last males to eclose also survived 24 days, guarding 3 females for 5 days and mating
twice (160 h). Females, when returned to their pupal tree (placed on the ground near
the tree), sought the tree and climbed upward. One female with an extraordinarily
distended abdomen climbed to over 5 m, ew down to a nearby maple and disap-
peared in the new foliage.
Predation and parasitism
In 2011, three newly-anchored larvae were preyed on as evidenced by remnant
anchored pygypod stumps. Possible predators were rodents, birds and spiders. From
2008-10, 14 of 94 pupae (15%) died. Twelve were eaten by spiders. A Johnson jumper,
Phidippus johnsoni group, and the star bellied orb weaver, Acanthepeira stellate,
were observed eating pupae and newly-eclosed adults, and leaving silk-wrapped
body parts attached to the tree. Some partially-eaten pupae survived to eclose albeit
with deformed elytrae and pronotums. From 2008-10, 17.4% (n = 132) of the larvae
and pupae did not progress to adults. A pupa that fell naked to the ground aer its
exuvia was eaten by an isopod woodlouse, Porcellio scaber, successfully eclosed as
a male 7 days later in the laboratory. Phorids, Antennatus apocephalus (Lloyd 1973,
Fig. 8) parasitised adults from 2008-11. Predator Photuris reies were not a signi-
cant threat to these early season reies as local Photuris were not active until May.
Mimics were abundant throughout the display season (Fig. 9).
Lampyrid Volume 2 (2012)
Dates and degree-days
From 2008-11, over half the nights in April had cold (< 12°C) or inclement weather
(e.g., Beaufort wind force 3+, rain, snow) unfavorable for ashing display. e larval
climb appeared related to relatively warm sunny days in February. Degree-days,
were useful for prediction of pupal and adult life stages and maximum display
days (Faust & Weston 2009). From 2008-11, both sexes combined (n = 78) averaged
eclosion on April 10 or aer 16 ± 5 days of pupation or 86 ± 16 mGDDC from pupae
to adult. e 18-year average for the occurrence of ash display behaviour at Knox
was 19 March – 24 April, mGDDC 119-234. Weather is extremely variable in April
e.g., 5 mGDDC/day in 2008 and 2009, and 7 mGDDC/day in 2010 and 2011, which
is seen in the varying pupal durations in calendar days, but more constant durations
in degree-days (Table 2). e higher elevation Sevier site had a later season with later
calendar dates but similar degree-day ranges.
Pupation trees
Pupation trees had certain traits: scaly or furrowed bark (10 ± 3 mm, n = 53), tall
(21-30 m), larger dbh trees (49 ± 14 cm, n = 73) with direct southern exposure to
the sun (located at forest margins or open gaps). Tulip poplars, hickories (Carya
pallida, C. cordiformis, C. glabra and C. tomentosa) and walnuts (Juglans nigra)
were used by 84% (n = 179) of the larvae; several oaks (Q. velutina, Q. alba, Q.
phellos and Q. bicolour) were used by 6%; three other tree species (pine: Pinus
virginiana; persimmon: Diospyros virginiana; and winged elm: Ulmus alata) were
used by 2.5%. Two 60 year-old Eastern Red Cedars (Juniperus virginiana) with
an abundance of stringy bark on smooth trunks, were the rst record of that tree
species as a pupation tree for P. borealis. ese cedars held eight pupae in 2011. Of
the 20 trees at Knox used in 2008, 18 were reused 2-4 seasons. In 2011, 4 of 18 trees
Table 2 Pyractomena borealis rst-sighting life-stage landmark date and degree-day range for
2008, 2009, 2010, 2011 for Knox, Jeerson, Morgan, Sevier Co., TN. One day = 3 yr. mean of 6.5
mGDDC in April. * 2012 larval climb and rst ash included. Average durations for total popula-
tion provided in tex t.
First Larval
First pupa First male First female First ash* Peak
Jan 22 -
Mar 1st
Mar 4th -
Mar 22nd
-Apr 8th
Apr 3rd -
Mar 19th -
Apr 12th
Apr 14th -
Apr 24th -
May 2nd
Lampyrid Volume 2 (2012)
were new. Since 2008, 62% of preferred marked trees (n = 74) had one pupa, 37%
had 2-7 pupae and 1% had 8-37 pupae. A lone sand hickory C. pallida surrounded
by choking kudzu Pueraria lobata sheltered 37 pupae in 2011, the most observed
pupae per one tree.
Trees used
e occurrence of both rey species on particular tree species is likely due to
preferential selection as these tree species are not the most abundant and comprise
less than 16% of the tree species in these forests (Oswalt 2007) (Fig. 3).
In congregating Asian rey species, GPS data on specic trees known to
attract mating congregations is being documented by the Malaysian Nature Society
(Mubah 2010), the Forest Research Institute Malaysia (FRIM) and others for the
rivers of Sarawak and Malaysia (Nada et al. 2009; Wong 2009; Jusoh et al. 2010;
Ohba & Wong 2012) and reects the growing understanding of the critical impor-
tance of preserving larval habitat and preferred trees, and educating the public of
this precious resource in both the East and West.
For the TN reies, which occupied rough-barked, large diameter trees whose
trunks received plenty of sunlight, tree selection may be based on availability of
trees that provide the most favorable conditions for the reies. Similar ndings
and conclusions were reported by Rooney and Lewis (2000) for E. corrusca adults
and Gentry (2003) for P. borealis larvae in FL; (in contrast, see Lloyd’s 1997 report of
preference for smooth-barked trees). e mass of the larger diameter trees combined
with south-facing, exposed trunks of the trees provide warmer spots needed for P.
borealis larvae early in the season when it is very cold, while deep bark furrows
provide shelter from rain, snow and wind and other weather extremes.
Calendar days and degree days
Because both species range over 2200 km north to south across the eastern US and
parts of Canada, use of degree-day ranges combined with traditional calendar date
ranges could be a more accurate method of determining local seasonal appearance
regardless of population location.
As evidenced each year including during the record breaking warmth in 2012,
rey life stage dates in calendar days can vary by 3-4 weeks (Faust & Weston
2009), but the degree day ranges remain more constant, even in the record breaking
warmth of 2012. Ellychnia corrusca appeared less inuenced by accumulated
degree days, though their coupling season was not reached until at ca. 100 mGDDC
accumulated, despite periods of warmth earlier in the season.
For P. borealis, both average calendar dates and mGDDs were useful in predicting
when certain life stages (pupal duration and eclosion by sex and peak display/court-
ship) would occur and when the season would end (Table 2). Increasing survey
Lampyrid Volume 2 (2012)
frequency during the predicted times for those life events helped capture impor-
tant information that may otherwise have been overlooked. Using degree-days as a
prediction tool could guide well-timed collecting eorts when limited by National
Park regulations and prevent over-collecting small local populations.
Predation and parasitism
Adults of E. corrusca suered early winter losses to probable rodent activity as
evidenced by the elytral remnants with bite marks found at the base of the colony
tree. In the milder winters of East TN, rodents such as mice and shrews are active
year-round and could pose a threat especially to E. corrusca near the ground. Rooney
and Lewis (2000) also found elytrae at the base of colony trees and suspected rodent
activity. e abrupt vertical emergence of the TN E. corrusca could help minimise
losses to rodent predation by allowing the reies to quickly ascend en masse up the
trees and away from the leaf litter. For P. borealis, the higher pupal losses to spider
predation and no (observed) predation of larvae raises the question as to why this
disparity exists. Larvae have chemical defenses (Lewis & Cratsley 2008) but it is
not known whether the same is true for the pupae. Early emergence in TN appears
to be an eective strategy by both rey species to avoid predation by Photuris
reies, which become active later and are important predators of P. borealis adults
elsewhere (Buschman 1974). Phorid parasitism of both species appeared to increase
as the season progressed (Fig. 8).
Additional discussion for Ellychnia corrusca
ese reies appear to overwinter as either adults or pupae somewhere other than
on exposed tree trunks and make a sudden initial appearance at the base of the
tree (and then climb up) during the rst relatively warm days of Jan-Feb, despite
more below-freezing weather being a certainty at this point of winter. In contrast,
the Massachusetts (MA) E. corrusca adults much further north, overwinter on the
trunks of their black oak colony trees (Rooney & Lewis 2000) beginning in early
fall. e more northerly Pennsylvania (Butler Co.) E. corrusca (pers. obs.) appears
to share the overwintering habit of the Massachusetts rey. In contrast, Hackett
et al. (1996) mention collecting E. corrusca in late winter 1983 “while they were
emerging from their overwintering sites at the base of a white oak tree in Belts-
ville, Maryland.” e more southern Maryland E. corrusca’s sudden appearance at
winter’s end more closely parallels the TN E. corrusca habit. Further studies are
needed to determine whether these regional dierences in emergence and overwin-
tering behaviours suggest distinct species, univoltine vs. semivoltine life cycles
perhaps inuenced by the milder climatic conditions in TN, or adaptive behaviours
governed by winter conditions and predation risks for dierent locales. Since E.
corrusca are a species complex (Fender 1970) instead of a distinct species, deter-
mining dierences in larval development, adult size variations, overwintering
habits and adult emergence times could help in species discrimination. No adults
Lampyrid Volume 2 (2012)
have yet been observed feeding in TN at sap ows or on owers as they have in the
more northern range. Perhaps the arboreal overwintering habit of the MA adults
requires more energy. Captive TN E. corrusca will however, feed from an apple slice,
sometimes choosing the apple over mating.
e clear drops of liquid that both sexes deposit from their terminal segment
when disturbed and/or climbing are in addition to the white beads of reexive
bleeding previously noted and described (Williams 1917; Rooney & Lewis 2000,
respectively). is clear liquid may be excreta, or a chemical cue promoting sexual
or social communication. e vertical upward migration on the days of initial
emergence (Fig. 7) suggests these reies are following one another up a particular
route of each tree and that something more than just the direction and heat from
the sun inuences this route
Further studies could focus on what actually occurs during the quiescent 4-6 wk
aer initial appearance but before ight and coupling. What is observed is that they
search the trunk or rest head down in the furrows and in the later weeks of quies-
cence, disappear into the canopy. e MA E. corrusca are known to be sexually
mature early in the season (Rooney & Lewis 2000) so perhaps unseen pre-ight
coupling is occurring in the canopy. Pre-ight mating, if true, would also reduce
risks associated with ight, e.g., exposure to the elements and predators, for these
weak yers.
Additional discussion for Pyractomena borealis
e ‘ckle’ behaviour of larvae in their constant search for a better pupation site
has also been observed previously (Lloyd 1997). TN larvae were especially mobile
when rst climbing in January and February and oen changed locations but not
trees before settling on a permanent pupation site usually by late February or early
March. Some larvae may disappear above the surveyed 3 m mark, where others
succumb to predation or the elements. Larvae pupated < 2 m high on the only two
smooth but stringy barked trees, yet the scaliest, roughest trees that are harder to
climb oen had pupation sites as high as 3m, a counter-intuitive nding. Further
study is needed to determine if TN larvae choose only furrowed trees for pupation,
or pupate out of sight above 3 m on rough and smooth-barked trees. Red and sugar
maples (Acer rubrum and Acer saccharum), which are smooth-barked species
until extreme age, are the most common trees in the local forests (Oswalt 2007)
comprising 17% of all stems >2.5cm dbh, yet P. borealis have not been found on any
maples (Fig. 3). Instead, 90% of pupae are found on poplars, oaks, hickories and
walnuts whose combined species comprise only 21% of the forest (Fig. 3).
Larvae avoid most of the forest oor vegetation that shades the lower tree trunks
by climbing up the trunk 1-2 m into the more direct warming rays of the sun.
Climbing higher has also been proposed as means to stay above high water levels
in ood-prone, low-lying forests in FL where P. borealis is found (Lloyd 1997). is
may also apply to the P. borealis that colonise trees in the alluvial forests in TN,
Lampyrid Volume 2 (2012)
though many of the preferred trees are located on high ground.
at the male larvae climb, attach, pupate and eclose earlier in the wild on
average than females concurs with ndings from a previous laboratory study
(Gentry 2003) where pupal durations were determined at constant temperatures.
Findings of Al-Saar et al. (1996) and Weston and Diaz (2005) indicate that varying
temperatures can actually enhance maturation in some insect species. Could it
be possible that the males obtain further developmental advantage by climbing
and pupating earlier in the season when the temperatures are more extreme and
variable? Is it possible that the guarding males provide some measure of thermal
protection to their tended female pupae thus inuencing female development (Figs.
10 & 11)?
Protandry gives the males much time to locate and guard potential mates, which
are oen coupled immediately aer the female’s eclosion. is diurnal mating
strategy, taking place during the relative warmth of the day, bypasses ash commu-
nication and may be an important adaptation to the nighttime conditions that are
oen unfavorable to courtship displays. Locating females without needing to y
or ash may increase males’ mating success and reserve their energy for photic
display on the few warm nights available when ight conditions are ideal. is dual
strategy approach ensures the males will have mating opportunities regardless of
the unpredictable late winter/early spring weather. While eclosing virgin females
had little control over mate choice, in rematings, depending on the situation and the
male, females displayed both acquiescent and eective avoidance behaviours. e
prolonged (up to 8 days) copulatory mate-guarding seen in P. borealis is common in
insects and is a male adaptation to avoid sperm competition (Tompkins & Simmons
In the past 19 years, the Knox reies have faced many challenges aecting their
once dark, forested rural habitat. e clearing of forested areas that the reies
inhabit and in which they display is a major threat, with 15% of the primary Knox
site lost in the past four years alone. While daytime mating incidentally precludes
E. corrusca from the negative eects of articial lighting, the nocturnal courtship
phase of P. borealis is not spared. Homes that have been built in these areas have
introduced light pollution and pollution from landscaping agrochemicals that
may be contributing to the apparent decline in rey populations. New shopping
centers, a football stadium and schools built less than 2 km away now illumi-
nate the night skies to the point that a ashlight is no longer necessary on some
nights while walking the rey survey trails. Habitat loss and light pollution are
also serious threats to Asian rey populations (Wong 2009; Nada et al. 2009) and
reies worldwide. Other imminent threats to the TN reies concern threats to
their colony trees. In addition to natural mortality each year due to high winds,
disease and lightning, the invasive kudzu vine Pueraria lobata threatens to smother
Lampyrid Volume 2 (2012)
many of the preferred and colony trees. e ousand Cankers disease (Geosmitha
spp.) that is spread by the walnut twig beetle, Pityophthorus juglandis (Coleop-
tera: Scolytidae) and is deadly to black walnuts, has recently been found infesting
walnuts in TN (M.T. Windham, pers. comm., August 2010).
Future studies of mechanisms governing the abrupt appearance and vertical
mass tracking up colony trees might provide new understanding of chemical
cues, early season adaptations and regional dierences in behaviour of the E.
corrusca complex. e described observations of male P. borealis daytime search
- and guard-behaviour of larva/pupal females, their copulation without courtship
dialogue early in the mating season, and their nocturnal habits and ash displays in
the latter half of the season, provide a rst insight into a dual mating strategy of a
lampyrid. With their preference for trees of a certain size, furrow depth, species and
specic location, both of these adaptable, hardy rey species nonetheless remain
vulnerable to natural and man-made changes to their environments. It is hoped
these studies will further the appreciation of the importance of individual trees
and habitat to the survival and health of larval, pupal and adult rey populations
and increase the understanding of these two species’ remarkable adaptations to the
habitat and climatic extremes of the early season.
My thanks to the hosts and sponsors of the 2010 International Firey Sympo-
sium held in Selangor, Malaysia for providing such a wonderful opportunity for
cooperation, collaboration, learning and sharing of information on the world of
reies. Special appreciation goes to the Forest Research Institute of Malaysia and
the Malaysian Nature Society. anks to Edgar Faust and Richard Evans for tree
details, to Ted Faust and Paul Weston for technical assistance, Haden Wesley and
Martie Wood for help in the eld and e Great Smoky Mountains National Park
(permit #FaustGRSM2009-SCI-0026) for access to two study sites.
A-S, Z., G, J.  A, J. (1996) Temperature and humidity
aecting development, survival and weight loss of the pupal stage of Drosophila
melanogaster and the inuence of alternating temperature on the larvae. Journal
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... Ellychnia corrusca is nicknamed the winter firefly because it has a winter-spring activity cycle, in contrast to the late spring-summer activity cycles of most other fireflies [2,10]. Ellychnia corrusca larvae in New England (North America) extend their larval stage across 2 years instead of one to ensure that they ingest ample calories, emerging in the late fall of their second year as adults and sexually maturing during the winter to start mating, which lasts until late spring [11]. Normally, adult fireflies do not need to feed because they only survive as adults for 2 to 4 weeks [2,4]. ...
... Normally, adult fireflies do not need to feed because they only survive as adults for 2 to 4 weeks [2,4]. However, E. corrusca firefly adults live Scott R. Smedley Deceased for~9 months, and in MA, USA, they ingest interstitial fluid and sap from maple trees to help them survive during the cold winters [11]. The Photuris versicolor species complex (hereafter: Ph. versicolor) contains > 17 firefly species that are differentiated by their male courtship lantern flashes but that cannot be identified from their morphology with confidence. ...
... Differences between the microbiomes of these two groups might be due to their different lifestyles. North American E. corrusca are active in winter and may feed on tree sap [10,11], and so E. corrusca microbial communities may therefore be acquired via the ingestion of these fluids. Ellychnia corrusca also live for~9 months. ...
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Although there are numerous studies of firefly mating flashes, lantern bioluminescence, and anti-predation lucibufagin metabolites, almost nothing is known about their microbiome. We therefore used 16S rRNA community amplicon sequencing to characterize the gut and body microbiomes of four North American firefly taxa: Ellychnia corrusca, the Photuris versicolor species complex, Pyractomena borealis, and Pyropyga decipiens. These firefly microbiomes all have very low species diversity, often dominated by a single species, and each firefly type has a characteristic microbiome. Although the microbiomes of male and female fireflies did not differ from each other, Ph. versicolor gut and body microbiomes did, with their gut microbiomes being enriched in Pseudomonas and Acinetobacter. Ellychnia corrusca egg and adult microbiomes were unique except for a single egg microbiome that shared a community type with E. corrusca adults, which could suggest microbial transmission from mother to offspring. Mollicutes that had been previously isolated from fireflies were common in our firefly microbiomes. These results set the stage for further research concerning the function and transmission of these bacterial symbionts.
... A surprising exception to this typical life cycle is seen in winter fireflies (Ellychnia corrusca), which overwinter as adults rather than as larvae. In this species complex, adults congregate in the fall and seek refuge in the deeply furrowed bark of large trees, sometimes using the same overwintering trees year after year (Faust 2012). These long-lived adults can survive for up to 10 months while overwintering on trees (Rooney and Lewis 2000). ...
... For example, eastern Pyractomena borealis fireflies, which pupate in tree bark furrows and emerge early in the year, often use the same large-diameter tulip poplars, hickories, and oaks year after year . Adults of the common day-active winter firefly (Ellychnia corrusca) also appear to prefer tree species with more deeply furrowed bark, and they can be found in loose colonies on these tree species as well (Faust 2012;Deyrup et al. 2017). Consider leaving some areas of your yard or property a little unkempt or more natural-the perimeter can work well for this if you want to keep wild areas to the edges. ...
... • The winter firefly (Ellychnia corrusca) overwinters as an adult in deeply furrowed bark of large trees (Faust 2012). Eastern Pyractomena borealis fireflies seek out bark furrows for pupation, often using the same large-diameter tulip poplars, hickories, and oaks year after year ). ...
Technical Report
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Fireflies are some of our most celebrated insects. They have immense cultural, biological, and economic importance and are important components of natural ecosystems. Their public appeal also makes them ideal flagship species for conservation. In this document, we outline the importance of fireflies and describe their life histories and threats to their populations. We then provide steps you can take to create and manage habitat, influence policies that benefit fireflies and their habitats, and raise awareness of this special group of beetles. We also provide a summary of the firefly genera found in the United States and Canada and a list of resources for learning more. Full text is available at
... Adult P. corruscus lack light organs and are diurnal. In contrast to familiar summer-active species, P. corruscus emerge as reproductively immature adults in the fall, overwinter on the sides of large-diameter trees, and mature their ovaries and testes in the spring for mating in the late spring/early summer (Rooney and Lewis 2000;Faust 2012;Hoagland 2022). Based on behavioral observations and comparison with related firefly species, males are likely the searching sex, flying toward perched calling females, landing nearby, and scrambling to approach (Lloyd 1972;Vencl and Carlson 1998;De Cock and Matthysen 2005). ...
... For field bioassays, because catch data were not normally distributed, the attractive effect of the pheromone relative to solvent controls was assessed using a one-sided Mann Whitney Wilcoxon test in R (R Core Team 2020). Locations were tested separately because there may be differences in phenology between the field sites due to latitude (Faust 2012). For laboratory assays, the difference in total time in contact with either control or experimental septa was assessed with paired Wilcoxon rank-sum tests. ...
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Firefly flashes are well-known visual signals used by these insects to find, identify, and choose mates. However, many firefly species have lost the ability to produce light as adults. These “unlighted” species generally lack developed adult light organs, are diurnal rather than nocturnal, and are believed to use volatile pheromones acting over a distance to locate mates. While cuticular hydrocarbons, which may function in mate recognition at close range, have been examined for a handful of the over 2000 extant firefly species, no volatile pheromone has ever been identified. In this study, using coupled gas chromatography - electroantennographic detection, we detected a single female-emitted compound that elicited antennal responses from wild-caught male winter fireflies, Photinus corruscus. The compound was identified as (1S)-exo-3-hydroxycamphor (hydroxycamphor). In field trials at two sites across the species’ eastern North American range, large numbers of male P. corruscus were attracted to synthesized hydroxycamphor, verifying its function as a volatile sex attractant pheromone. Males spent more time in contact with lures treated with synthesized hydroxycamphor than those treated with solvent only in laboratory two-choice assays. Further, using single sensillum recordings, we characterized a pheromone-sensitive odorant receptor neuron in a specific olfactory sensillum on male P. corruscus antennae and demonstrated its sensitivity to hydroxycamphor. Thus, this study has identified the first volatile pheromone and its corresponding sensory neuron for any firefly species, and provides a tool for monitoring P. corruscus populations for conservation and further inquiry into the chemical and cellular bases for sexual communication among fireflies.
... In temperate latitudes, fireflies seem to occur almost exclusively during the spring and summer seasons (e.g. Majka 2012;Faust 2017), with few exceptions (Faust 2012). However, in the tropics, fireflies have been found to have species-specific temporal occurrences that may vary in duration (i.e. ...
... Firefly species with adults being active both during the day and at night are rare, although this phenomenon could have been overlooked. In the best documented case of a firefly species active in both periods, the males of the winter firefly Pyractomena borealis adopt two strategies: in the early season, they seek out females during the day, bypassing courtship with flashing at night; in the late season, they switch to a nocturnal habit and use light signals to communicate with females (Faust 2012). The seasonal variation of the daily period of activity in Photuris elliptica and Pyrogaster nigrolineatus remains unknown. ...
Fireflies (Coleoptera: Lampyridae) are charismatic insects that have been fruitful model systems in biotechnology. However, lack of information about firefly taxonomy and ecology renders species identification a hard task, especially in the Neotropical region, where fireflies are most diverse. A major gap in the literature on Neotropical fireflies is the lack of knowledge on species’ ecological niches and habitats, which are fundamental aspects for understanding their biology. Here, we provide an annotated checklist of the firefly fauna of the Serra dos Órgãos mountain range (Rio de Janeiro State, Southeastern Brazil), with information on the natural history of each species. We assembled data in three ways: monthly sampling with Malaise traps and active search along an elevational transect from 130 to 2,170 m, over 2 years (2014–2016), extensive field observations, and extracted from historical species records for the Serra dos Órgãos from museum specimens in key collections in Brazil and Europe. We provide a taxonomic key to the genera recorded in the region, and a differential diagnosis for each species, highlighting key references for each taxon. We report 58 species representing 21 genera, making the Serra dos Órgãos one of the richest firefly hotspots on Earth. Most species are restricted to one or two habitat types and/or just one of the regional seasons (warm or cool), and many were only collected either by malaise traps or active search, underlining the importance of sampling different habitat types and seasons, and using different sampling methods when surveying fireflies. Out of the 51 species observed in the field, 49 were active either during the day or the night, although two species – Photuris elliptica and Pyrogaster nigrolineatus – were active in both periods, which is rare in fireflies.
... This flash exchange -known as a courtship dialog -continues until the male locates and mounts the female, and is characteristic of many firefly genera including Luciola, Aquatica, Pyractomena, and Photinus (Stanger- Hall and Lloyd, 2015). Females in the genus Photuris (Souto et al., 2019) are specialist predators of other flashing fireflies, luring patrolling males by mimicking the response flashes of conspecific females (Lloyd, 1980), hawking them from the air (Lloyd and Wing, 1983), and stealing them from spider webs (Faust, 2012). In a few species, such as Photinus carolinus, Photinus knulli, and Photuris frontalis in the United States (Sarfati et al., 2021(Sarfati et al., , 2022, Photinus palaciosi in Mexico, and Luciola sp. in eastern Europe (Baldaccini et al., 1969), flying males synchronize their courtship advertisements when they reach sufficient densities. ...
Full-text available
Bioluminescent insects have been the subject of scientific interest and popular wonder for millennia. But in the 21st century, the fireflies, click beetles, and cave glow-worms that brighten our nights are threatened by an unprecedented competitor: anthropogenic light pollution. Artificial lights can obscure the light-based signals on which these and other bioluminescent organisms rely to court mates, deter predators, and attract prey. In the following review we summarize a recent influx of research into the behavioral consequences of artificial light at night for firefly beetles (Coleoptera: Lampyridae), which we organize into four distinct courtship signaling systems. We conclude by highlighting several opportunities for further research to advance this emerging field and by offering a set of up-to-date lighting recommendations that can help land managers and other stakeholders balance public safety and ecological sustainability.
... We conducted the field portion of this study at Massachusetts Audubon's Habitat Sanctuary in Belmont, Massachusetts, USA (42.402184°N, 71.187205°W), a site we have used previously to investigate the behavior and life history of E. corrusca (Rooney & Lewis 2000). Widely distributed across the eastern US, this diurnal firefly has a unique phenology (Rooney & Lewis 2000;Faust 2012;Deyrup et al. 2017). In New England, the non-luminescent E. corrusca adults eclose in early fall, spend the winter exposed on tree trunks, then mate in early spring. ...
While fireflies are most renowned for their bioluminescent signals, several species also have been shown to possess UV and blue-induced fluorescence. In a field study, we used a sentinel prey approach to investigate whether this trait might serve as an aposematic function in chemically defended fireflies. We also examined and provide the first detailed description of green and red patterns of blue light-induced fluorescence in the adults and larvae of Ellychnia corrusca (L.) (Coleoptera: Lampyridae). We placed artificial prey models with and without fluorescent markings in a forested area of Belmont, Massachusetts, USA, where they were exposed to natural predators. Clay models were styled to resemble Photinus fireflies, the most common firefly genus in this area, and a close relative of Ellychnia. The field study was conducted from Oct through May in Belmont, Massachusetts; this is the period when adults of the diurnal non-luminescent firefly E. corrusca are overwintering on trees. We observed considerable seasonal variation in attack rates, and marks on the clay models indicated they were attacked by avian, mammal, and arthropod predators. However, no difference was seen between fluorescent and non-fluorescent models in predator attack rates. This suggests that fireflies' fluorescent markings do not serve as an aposematic signal, at least for the predator guild active in New England woodlands.
... Using the "modified corn growing degree day or 86/50 Fahrenheit" (mGDD) with 1 March start date formula used in prior studies (Faust and Weston 2009;Faust 2010Faust , 2012Faust , 2017Faust and Forrest 2017), P. walldoxeyi was not yet active at ca. 589 mGDD on 5 May 2018 at the type locality, yet had appeared in low to moderate numbers by 12 May at 739 mGGD and were still present in declining For northern Mississippi, the first appearance of P. walldoxeyi in early May represents the earlier phase of the predictable seasonal progression of the local firefly species assemblage. Pyractomena borealis (Randall) (L. ...
Photuris walldoxeyi Faust, new species, is described as a member of the Photuris versicolor species-group. It can be distinguished by its complex, unique flash-train + glow (FT+G) male courtship flash pattern, one of the most easily recognizable displays in North America. The primary FT+G male courtship pattern consists of a flash-train of 4-9 bright, quick, rhythmic pulses given every 0.4 seconds followed by a prolonged diffuse glow (ca. 1 second) of medium intensity at 17° C while the male hovers flashing near the tips of the branches of bald cypress and swamp vegetation growing in water. The primary FT+G is often followed immediately by a 2-pulsed secondary FT+G that effectively increases the male's visibility. Photuris walldoxeyi has thus far been confirmed in cypress and tupelo-cypress swamps of Mississippi, west Tennessee, and Indiana. During its month-long flight from May to mid-June, peak numbers are present ca. 900-1200 modified growing degree days.
Full-text available
Firefly flashes are well-known visual signals used by these insects to find, identify, and choose mates. However, many firefly species have lost the ability to produce light as adults. These unlighted species generally lack developed adult light organs, are diurnal rather than nocturnal, and are believed to use volatile pheromones acting over a distance to locate mates. While cuticular hydrocarbons, which may function in mate recognition at close range, have been examined for a handful of the over 2000 extant firefly species, no volatile pheromone has ever been identified. In this study, using coupled gas chromatography - electroantennographic detection, we detected a single female-emitted compound that elicited antennal responses from wild-caught male winter fireflies, Photinus corrusca . The compound was identified as (1S)- exo -3-hydroxycamphor (hydroxycamphor). In field trials at two sites across the species eastern North American range, large numbers of male P. corrusca were attracted to synthesized hydroxycamphor, verifying its function as a volatile sex attractant pheromone. Males spent more time in contact with lures treated with synthesized hydroxycamphor than those treated with solvent only in laboratory two-choice assays. Further, using single sensillum recordings, we characterized a pheromone-sensitive odorant receptor neuron in a specific olfactory sensillum on male P. corrusca antennae and demonstrated its sensitivity to hydroxycamphor. Thus, this study has identified the first volatile pheromone and its corresponding sensory neuron for any firefly species, and provides a tool for monitoring P. corrusca populations for conservation, and further inquiry into the chemical and cellular bases for sexual communication among fireflies.
Full-text available
Background Fireflies (Coleoptera: Lampyridae) are commonly recognized by adult traits, such as a soft exoskeleton, lanterns and associated glow and flash patterns, but their larval stage is far less appreciated. However, fireflies spend most of their lives as larvae, and adults of most species rely solely on resources previously obtained. Therefore, studying the immature stages is imperative towards a comprehensive understanding of fireflies. This paper reviews and indicates key gaps in the biology of firefly larvae based on available literature. Methodology We reviewed the literature on firefly larvae to identify key issues and important taxonomic, geographic, and subject biases and gaps. Results We found 376 papers that included information on firefly larvae. Only 139 species in 47 genera across eight of eleven lampyrid subfamilies have been studied during larval stages. These numbers reveal a staggering gap, since 94% of species and over half of the genera of fireflies were never studied in a crucial stage of their life cycle. Most studies on firefly larvae focus on two subfamilies (Luciolinae and Lampyrinae) in four zoogeographic regions (Sino-Japanese, Oriental, Nearctic, and Palearctic), whereas the other subfamilies and regions remain largely unstudied. These studies mainly dealt with morphology and behavior, other subjects remaining greatly understudied by comparison, including habitats, life cycle, physiology and interactions. Conclusions Together, these literature biases and gaps highlight how little is known about firefly larvae, and warmly invite basic and applied research, in the field and in the lab, to overcome these limitations and improve our understanding of firefly biology to better preserve them.
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Example insect natural history data (PDF document)
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Pyrrhalta viburni (Paykull), a new landscape pest in the United States, is univoltine and overwinters in the egg stage, feeding as larvae and adults on the foliage of plants in the genus Viburnum. Experiments were conducted to determine chilling requirements for completion and breaking of diapause and development rate of larvae at different temperatures in the laboratory. Egg hatch required at least 4 mo for induction of diapause, and &ap;200 growing degree-days (GDD) at a calculated base temperature of 8.2°C for breaking of diapause in the laboratory. Development of all larval instars was significantly faster at 22 than at 17°C and faster at 27 than at 22°C, but 27°C appeared to be nearing the upper developmental threshold, because mortality was uniformly high, and deformities of adults were observed. We confirmed that larvae pass through three instars and found that head capsule width increased geometrically with instar. We calculated the larval developmental threshold to be &ap;11.7°C at constant temperatures in the laboratory. Development under ambient conditions, where temperatures fluctuate, was considerably faster than that observed in the laboratory at constant temperatures. Thus, rearing experiments conducted in the laboratory at constant temperatures provide useful estimates of threshold temperatures but unrealistic estimates of GDD thresholds for P. viburni and other insects whose developmental rate differs between constant and fluctuating temperatures.
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Population dynamics and reproductive activity were examined in a Massachusetts population of the common diurnal firefly, Ellychnia corrusca. Although closely related to nocturnal beetles in the genus Photinus, Ellychnia lack adult light organs. A mark-recapture study of overwintering adults demonstrated low winter mortality and supported the hypothesis that adults overwinter for a single year. By dissecting males and females sampled throughout late winter and spring, it was found that adults become reproductively active in early March, when male seminal vesicles first contained sperm and female ovaries first contained mature oocytes. Both sexes mated multiply during the approximately six-week mating season (early April through mid-May), and copulations lasted up to 28 h. Adults collected in fall had higher abdominal fat body volumes than those collected in spring, and females contained more fat body than males. The life history and mating behavior off. corrusca are discussed in comparison to Photinus fireflies.
The firefly Pyractomena lucifera (Melsh.) occurs in fresh water marshes throughout the eastern half of temperate North America. Larvae captured prey both above and below the water surface and dragged it above water to feed. Prey records included: snails (n = 38) (Gastropoda: Pulmonata), freshwater limpets (n = 5) (Gastropoda: Ancylidae), a jumping spider (Salticidae), a damselfly nymph (Odonata) and a leech (Annelida: Hirudinea). Cryptically pigmented pupae were found on emergent vegetation and did not glow when probed. The pupal stage lasted 6.8 days for males and 6.4 days for females. At dusk males flew over the vegetation emitting single flashes (0.2 sec long, 27°C) at 2.9-5.1 sec intervals (17-24°C). Females answered male flashes with single flashes (ca. 1 sec long) at delays of 0.7-1.5 sec (17-27°C). Mated females seldom answered male flashes. Females oviposited when they were 5-6 days old, but 2-4 days after mating. They laid 30-194 eggs (mean = 102) and the number of eggs laid correlated with pupal weight of the female (correlation coefficient = 0.82). Eggs measured 0.8 mm and hatched in 15 days. They became faintly luminescent 2 or 3 days after oviposition and remained luminescent until they hatched. Six predators of this firefly were recorded: wolf spiders (n = 2) (Lycosidae), an orb weaver spider (Argiopidae), a harvestman (Phalangida), a giant water bug (Belostomatidae) and a tree frog (Hyla sp.). /// La luciérnaga Pyractomena lucifera (Melsh.) ocurre en pantanos de agua dulce a través de la mitad éste de la zona templada de Norteamerica. Las larvas capturan la presa arriba y debajo de la superficie del agua y la llevan arriba del agua para comérsela. Se han reportados como presas: caracoles (n = 38) (Gastropoda: Pulmonata), "limpets" de agua dulce (n = 5) Gastropoda: Ancylidae), una araña saltadora (Salticidae), una ninfa "damselfly" (Odonata) y una sanguijuela (Annelida: Hirudinea). En vegetación nueva se encontraron pupas crípticamente pigmentadas y no brillaron cuando se tocaron. El estado pupario duró 6.8 días en los machos y 6.4 días en las hembras. Al anochecer, los machos volaron sobre la vegetación emitiendo destellos individuales (0.2 segundos, 27° C) a intervalos de 2.9 a 5.1 segundos (17-24°C). Las hembras le contestaron a los machos con destellos individuales (aprox. 1 segundo de duración) con demoras de 0.7-1.5 segundos (17-27° C). Hembras que habían copulado, rara vez le contestaban al destello de los machos. Las hembras pusieron los huevos cuando eran de 5-6 días de edad, pero después de 2-4 días de haber copulado. Ellas pusieron de 30-194 huevos (promedio = 102) y el número de los huevos puestos está correlacionado con el peso de las pupas de las hembras (coeficiente de correlación = 0.82). Los huevos midieron 0.8mm y nacieron en 15 días. Dos o tres días después de puestos los huevos, eran vagamente luminicentes y se mantuvieron así hasta que nacieron. Se han reportado 6 rapazes de esta luciérnaga: arañas "wolf" (n = 2) (Lycosidae), una araña "orb weaver" (Argiopidae), un "harvestman" (Phalangida), un "water bug" gigante (Belostomatidae) y una rana de árbol (Hyla sp.).
Research at institutions of higher education could be restored to at least a shadow of its original role through publication in a manner appropriate for immediate classroom use, with questions that pique and direct the interests and activities of students. Studies on basic natural history may be good candidates for such publication and an example is drawn from fireflies: Two woodland species show directional orientation in their pupation sites on the trunks of trees; one uses southerly exposure and the other occurs on the north side of smaller trees, and much lower on the trunks. These contrasting positions have different thermal consequences, as demonstrated with a physical model, which possibly have a role in reducing interspecific sexual contact or prey competition. /// La investigación en instituciones de educación avanzada podría ser restaurada parcialmente a su rol original a traves de publicaciones, de manera tal que las mismas puedan ser usadas para enseñar, con preguntas que atraigan el interés de estudiantes y que se relacionen con sus actividades. Los estudios de historia natural básica pueden ser buenos candidatos para ese tipo de publicaciones, y un ejemplo del mismo se puede obtener con luciérnagas: Dos especies de luciérnagas muestran diferencias en la ubicación de sus pupas en los troncos de los árboles; una especie las ubica expuestas hacia el sur y la otra usa el lado norte de árboles mas pequeños y en la zona mas baja del tronco. Estas posiciones contrastantes tienen diferentes consecuencias térmicas, como se demuestra con un modelo físico, las cuales podrían tener un papel en reducir el contacto sexual o la competencia por alimento entre las dos especies.
Landscape-driven microclimates in mountainous terrain pose significant obstacles to predicting the response of organisms to atmospheric warming, but few if any studies have documented the extent of such finescale variation over large regions. This paper demonstrates that ground-level temperature regimes in Great Smoky Mountains National Park (Tennessee and North Carolina) vary considerably over fine spatial scales and are only partially linked to synoptic weather patterns and environmental lapse rates. A 120-sensor network deployed across two watersheds in 2005-06 exhibited finescale (,1000-m extent) temperature differences of over 2*C for daily minima and over 4*C for daily maxima. Landscape controls over minimum temperatures were associated with finescale patterns of soil moisture content, and maximum temperatures were associated with finescale insolation differences caused by topographic exposure and vegetation cover. By linking the sensor array data to 10 regional weather stations and topographic variables describing site radiation load and moisture content, multilevel spatial models of 30-m resolution were constructed to map daily temperatures across the 2090-km2 park, validated with an independent 50-sensor network. Maps reveal that different landscape positions do not maintain relative differences in temperature regimes across seasons. Near-stream locations are warmer in the winter and cooler in the summer, and sites of low elevation more closely track synoptic weather patterns than do wetter high-elevation sites. This study suggests a strong interplay between near-ground heat and water balances and indicates that the influence of past and future shifts in regional temperatures on the park's biota may be buffered by soil moisture surfeits from high regional rainfall.